An introduction to Teleutomyrmex.....

Strongylognathus Mayr, 1853d.

Strongylognathus [Myrmicinae:Tetramoriini].
- Strongylognathus Mayr, 1853d: 389.Type-species: Eciton testaceum, by monotypy. Replacement name for Myrmus Schenck, 1853: 188. [Junior homonym of Myrmus Hahn, 1832: 81 (Hemiptera).].

Myrmus [junior homonym, see Strongylognathus].
- Myrmus Schenck, 1853: 188.Type-species: Myrmus emarginatus (junior synonym of Eciton testaceum), by monotypy. [Junior homonym of Myrmus Hahn, 1832: 81 (Hemiptera).].
- Strongylognathus Mayr, 1853d: 389, replacement name for Myrmus Schenck.

Myrmus Schenck, 1853.

Myrmus [junior homonym, see Strongylognathus].
- Myrmus Schenck, 1853: 188.Type-species: Myrmus emarginatus (junior synonym of Eciton testaceum), by monotypy. [Junior homonym of Myrmus Hahn, 1832: 81 (Hemiptera).].
- Strongylognathus Mayr, 1853d: 389, replacement name for Myrmus Schenck.

Some know why I putted this one here..............

Now, let me explain. It was shown by Ward et al. 2015 ("2014") that Teleutomyrmex and Anergates should be synonymized under Tetramorium but, surprisingly, they kept Strongylognathus out of it. It should be a synonym of Tetramorium but the name should not be Strongylognathus but Tetramorium ("better name'). But if you do it, do it good and synonymize it like it should... so: Strongylognathus.

Tetramorium Mayr, 1855.

Tetramorium [Myrmicinae: Tetramoriini].
- Tetramorium Mayr, 1855: 423. Type-species: Formica caespitum, by subsequent designation of Girard, 1879: 1016.

Extreme, Workerless Inquiline ants.

Ants: Probably one of the most important animal-groups on Earth. Together with wasps and bees they play some of the most important roles in ecosystems around the world. Without these three groups of insects, most animal- and plant-species will become extinct on this little globe in space in the near future. So remember them well: Ants - Formicidae, Wasps - Vespidae and Bees (including digger-wasps) – Apidae. And if you want to know much about them, be prepared to read a lot! Almost every conceivably down-to-earth lifestyle you can think of, somewhere one or other ant lives like that, and for the airborne lifestyles, go to the wasps and bees! Have a lifetime of fun to discover all these animals! And for the most specialized animals on earth, you must go to the ants and discover the species in the genera Teleutomyrmex and Anergates and a few other, mostly related, species!

The extreme, workerless inquilines.

The genus Teleutomyrmex was described in 1950 by Heinrich Kutter, based on ants discovered in Saas-Fee (a small town in the canton Wallis, Switzerland.) in 1949 and 1950. The species was named T. schneideri. Later, the species was also found in the French Alps and Pyrenees, the Spanish Pyrenees, in Turkmenistan and in a nearby place in the Swiss Alps. The type-meadow in Saas-Fee was destroyed between 1950 and 1971. A second species, T. kutteri, was described in 1990 by Alberto Tinaut based on animals from the Sierra Nevada, Spain and the third and forth species, from 2017, are from Bulgaria and Turkey, T. buschingeri and T. seiferti.

The ants from the genus Teleutomyrmex are the most specialized ants on Earth. They are extreme, workerless inquilines that became ectoparasites. Let me explain.

- Parasite: An animal that is dependent on another species to survive. This dependence is temporary (during a certain period of its life.) or permanent.
- Social parasites: Social animals (like ants!) that are dependent on other social animals to survive. This can be temporary (during colony-foundation.) or permanent.
- Inquilines: Permanent social parasites among ants are also called inquilines.
- Workerless inquilines: The worker-caste, not needed by the inquilines, has disappeared. Only females/queens and males exist.
- Extreme, workerless inquilines: The females/queens and males have undergone some important morphological changes. Through these changes the ants are more adapted to their specialized way of life but, at the same time, they make sure that the ants can’t survive without their host. Some examples are: reduction of the mouthparts, development of appeasement-glands, over-development of the reproduction-organs, becoming weak and “soft”, males that become pupoid (show characteristic modifications that makes the male look “like a pupa”, e.g. yellowish color, downward curved gaster, big external genital plates,...),…
- Ectoparasites: Parasites that need to be carried around by their hosts. They are not capable to or have great difficulty with walking very short distances.

Edward Osborne Wilson, in 1971, wrote down a list with almost all the characters that determine social parasites and, in 1990, completed the list together with Berthold K. Hölldobler. In it 41 characters are listed that extreme, workerless inquilines can have. Not all those inquilines have all the characters but they have most of them. The ants of the genus Teleutomyrmex display 36 characters of the list but have also a few adaptations that are special for their ectoparasitic lifestyle.

Only around fourteen ant species are known that are extreme, workerless inquilines and four of them, the Teleutomyrmex species, have become ectoparasites. Teleutomyrmex females/queens have, for example, unique morphological adaptations like the dorsoventrally compressed gaster that easily fit around the gaster of the host-queen. Also, the queens and males of Teleutomyrmex-species have the terminal tarsal segments of their legs adapted/modified to be able to grip firmly the body of queens and workers of the host species and are so almost completely unable to walk alone.

One remarkable fact is that the males of Teleutomyrmex still have rudimentary, unusable wings while the males of the other extreme, workerless inquilines have lost the wings completely. In all its other characteristics it is further evolved compared with the other species. Strange but true!

A peculiar taxonomic fact: The genera Anergates and Teleutomyrmex and the host-genus Tetramorium are closely related ant-genera belonging to the tribe (group of closely related genera.) Tetramoriini (now the Crematogastrini). This tribe is part of the subfamily Myrmicinae which also includes the genus Pheidole (in the tribe Pheidolini, now the Attini!). Only Nylanderia belongs to a different subfamily, the Formicinae. Most of the known social parasitic ants belong to these two subfamilies…

Some taxonomic problems in this group of specialized ants.

In 1950, William Steel Creighton placed Anergates friedlandi as a synonym of A. atratulus. Although he recognized certain morphological differences between the two species in 1934 when he described A. friedlandi, he based his 1950 decision on the speculation of William L. Brown Jr. that the host-species of both inquilines was the same (yes) and that the North American population wasn’t native to that continent (maybe). So, no morphological data but host-species distribution was used to establish the synonymy. A few myrmecologists still question the decision and wait for the ongoing genetic comparison of European and North American Anergates-samples. Now it is clear that the North American form is an introduced population of A. atratulus!

The last few years some myrmecologists (like Alfred Buschinger) think that the morphological differences between Teleutomyrmex schneideri and T. kutteri are very minimal and question if T. kutteri should be placed as a synonym of T. schneideri. Most still think both deserve species status (clear morphological differences between the queens and the males of both species!) and, for the moment, both names stay as species names on record. Now, with the description of two more species, things are getting clearer in this genus.

Paratrechina and related genera were reviewed on genera-level in 2010 and one of the results was the division of Paratrechina into a few related genera, including Nylanderia. Some of these genera and part of the genus Nylanderia are recently revised and Nylanderia from North America was revised also but without the inquilines.

Tetramorium and related parasitic genera underwent a very thorough genetic phylogenetic study by Matthias Sanetra and Alfred Buschinger in 2000. When you read the paper only two possibilities exists. The first one: Tetramorium, Anergates and Teleutomyrmex should be considered to be synonyms of Strongylognathus and all the species together form one big genus (as Ward et al., 2015 ("2014"), say!). The other one: Tetramorium should be divided in at least eight different genera, all standing together with the parasitic ones in one compact tribe. The authors of the article still can’t follow either of the possibilities. Who will take a decision?

In 1951 Walter Linsenmaier made a crude black-and-white drawing of three Teleutomyrmex schneideri females riding on the back of a Tetramorium-queen. In front of this group, a Tetramorium-worker stands alongside her queen. This drawing was published in the 1951-article that Robert Stumper wrote about the life-style of the Teleutomyrmex schneideri-queen.

In 1951 a color-version of this drawing was also made by Walter Linsenmaier for an article in "Du: kulturelle Monatsschrift" by Heinrich Kutter, “Von dufttäuschenden Mörderinnen und berittener Königin.”. Heinrich Kutter used it again in 1968 as the front-plate of his little book about social parasitic ants of Switzerland. Later Walter Linsenmaier used it in his general review-book of insects of the world, published in 1973/4.

In 1977 a better version of the black-and-white drawing appeared in Heinrich Kutter’s determination-book for Swiss ants. This version is the most-copied drawing when Teleutomyrmex is depicted in an article or a book.

And on June 12, 2011 a colored version of the black-and-white/color-version drawing appeared as a tattoo on the back of my right shoulder… 60 years after its first publication!

And again with the article the topic started with, but this time on the pages of the magazine itself:

https://www.myrmecologicalnews…tegory&id=1493&Itemid=407 or https://www.myrmecologicalnews…_printable.pdf&format=raw

Amonio David Cuesta-Segura, Fede García García, Chema Catarineu, Sergio García-Tejero & Xavier Espadaler (2018): ACTUALIZACIÓN DE LA DISTRIBUCIÓN Y HOSPEDADORES DE LA HORMIGA PARÁSITA TELEUTOMYRMEX SCHNEIDERI KUTTER, 1950 EN LA PENÍNSULA IBÉRICA (HYMENOPTERA: FORMICIDAE).
Boletín de la Sociedad Entomológica Aragonesa (S.E.A.), nº 63 (31/12/2018): 235–239.

https://www.researchgate.net/p…ca_Hymenoptera_Formicidae

(Thank you Merkur!)

Philippe WEGNEZ, David IGNACE, Els LOMMELEN, Maximilien HARDY, Johan BOGAERT & Carin NILSSON (2015) : Redécouverte de Teleutomyrmex schneideri Kutter, 1950 dans les Alpes françaises (Hymenoptera: Formicidae). - Bulletin de la Société royale belge d’Entomologie/Bulletin van de Koninklijke Belgische Vereniging voor Entomologie, 151: 52-57.

http://www.antwiki.org/wiki/im…Alpes_fran%C3%A7aises.pdf

Cuesta-Segura, A.D., Espadaler, X., Garcia Garcia, F., 2017, “Hormigas de los brezales de Calluna cantábricos (NO España) (Hymenoptera: Formicidae). [Ants of the Cantabrian Calluna-heathlands (NW Spain) (Hymenoptera: Formicidae).]” Iberomyrmex, vol. 9, p. 25-43.

http://www.mirmiberica.org/node/626

From: AntWiki - Where Ant Biologists Share Their Knowledge.

Hölldobler, B. and Wilson, E. O. 1990. The Ants. Cambridge, Mass. Harvard University Press. Text used with permission of the authors.

From Chapter 12:

The “ultimate” social parasite.

There is no better way to begin a survey of the social symbioses than by considering the most extreme example known, that of the “ultimate” parasitic ant Teleutomyrmex schneideri. This remarkable species was discovered by Heinrich Kutter (1950a) at Saas-Fee, in an isolated valley of the Swiss Alps near Zermatt. Its behavior has been studied by Stumper (1950) and Kutter (1969), its neuroanatomy by Brun (1952), and its general anatomy and histology by Gösswald (1953). A second population has been reported from near Briançon in the French Alps by Collingwood (1956), a third in the French Pyrenees by Buschinger (1987c), and still others in the Spanish Sierra Nevada by Tinaut Ranera (1981). Appropriately, the name Teleutomyrmex means “final ant.”
The populations of Teleutomyrmex schneideri, like those of most workerless parasitic ant species (Wilson, 1963), are small and isolated. The Swiss population appears to be limited to the eastern slope of the Saas Valley, in juniper-Arctostaphylos woodland ranging from 1,800 to 2,300 m in elevation. The ground is covered by thick leaf litter and sprinkled with rocks of various sizes, providing, in short, an ideal environment for ants The ant fauna is of a typically boreal European complexion, comprising the following free-living species listed in the order of their abundance (Stumper, 1950): Formica fusca, Formica lugubris, Tetramorium caespitum, Leptothorax acervorum, Leptothorax tuberum, Camponotus ligniperda, Myrmica lobicornis, Myrmixa sulcinodis, Camponotus herculeanus, Formica sanguinea, Formica rufibarbis, Formica pressilabris, and Manica rubida. For some unexplained reason this little assemblage is extremely prone to social parasitism. Formica sanguinea is a facultative slavemaking species, preying on the other species of Formica. Doronomyrmex pacis, a workerless parasite living with Leptothorax acervorum, was discovered by Kutter as a genus new to science in the Saas-Fee forest in 1945. In addition, Kutter and Stumper found Epimyrma stumperi in nests of Leptothorax tuberum, as well as two parasitic Leptothorax, goesswaldi and kutteri, in nests of Leptothorax acervorum (Kutter, 1969).
Teleutomyrmex schneideri is a parasite of Tetramorium caespitum and Tetramorium impurum. Like so many other social parasites, it is phylogenetically closer to its host than to any of the other members of the ant fauna to which it belongs. In fact, it may have been derived directly from a temporarily free-living offshoot of this species, since Tetramorium caespitum and Tetramorium impurum (the host species at Briançon and in the Pyrenees) are the only nonparasitic tetramoriines known to exist at the present time through most of central Europe. It is difficult to conceive of a stage of social parasitism more advanced than that actually reached by Teleutomyrmex schneideri. The species occurs only in the nests of its hosts. It lacks a worker caste, and the queens contribute in no visibly productive way to the economy of the host colonies. The queens are tiny compared with most ants, especially other tetramoriines; they average only about 2.5 mm in total length. They are unique among all known social insects in being ectoparasitic. In other words, they spend much of their time riding on the backs of their hosts (Figure 12-1). The Teleutomyrmex queens display several striking morphological features that are correlated with this peculiar habit. The ventral surface of the gaster (the large terminal part of the body) is strongly concave, permitting the parasites to press their bodies close to those of their hosts. The tarsal claws and arolia are unusually large, permitting the parasites to secure a strong grip on the smooth chitinous body surface of the hosts. The queens have a marked tendency to grasp objects. Given a choice, they will position themselves on the top of the body of the host queen, either on the thorax or the abdomen. Deprived of the nest queen, they will then seize a virgin Tetramorium queen, or a worker, or a pupa, or even a dead queen or worker. Stumper observed a case in which six to eight Teleutomyrmex queens simultaneously grasped one Tetramorium queen, completely immobilizing her. The mode of feeding of the Teleutomyrmex is not known with certainty. The adults are evidently either fed by the host workers through direct regurgitation or else share in the liquid regurgitated to the host queen. In any case, they are almost completely inactive most of the time. The Teleutomyrmex adults, especially the older queens, are highly attractive to the host workers, who lick them frequently. According to Gösswald, large numbers of unicellular glands are located just under the cuticle of the thorax, pedicel, and abdomen of the queens; these are associated with glandular hairs and are believed to be the source of a special attractant for the host workers. The abdomens of older Teleutomyrmex queens become swollen with fat body and ovarioles, as is shown in Figure 12-1. This physogastry is made possible by the fact that the intersegmental membranes are thicker and more sclerotized than is usually the case in ant queens and can therefore be stretched more. Also, the abdominal sclerites themselves are widely overlapping in the virgin queen, so that the abdomen can be distended to an unusual degree before the sclerites are pulled apart. The ovarioles increase enormously in length, discard their initial orientation, and infiltrate the entire abdomen and even the postpetiolar cavity.
From one to several physogastric queens are found in each parasitized nest, usually riding on the back of the host queen. Each lays an average of one egg every thirty seconds. The infested Tetramorium colonies are typically smaller than uninfested ones, but they still contain up to several thousand workers. The Tetramorium queens also lay eggs, and these are capable of developing into either workers or sexual forms (Buschinger, personal communication). Consequently the brood of a parasitized colony consists typically of eggs, larvae, and pupae of Teleutomyrmex queens and males mixed with those of Tetramorium workers.
The bodies of the Teleutomyrmex queens bear the mark of extensive morphological degeneration correlated with their loss of social functions. The labial and postpharyngeal glands are reduced, and the maxillary and metapleural glands are completely absent. The mandibular glands, on the other hand, are apparently normal. In addition, the queens possess a tibial gland, the function of which is unknown. The integument is thin and less pigmented and sculptured in comparison with that of Tetramorium; as a result of these reductions the queens are shining brown, an appearance that contrasts with the opaque blackish brown of their hosts. The sting and poison apparatus are reduced; the mandibles are so degenerate that the parasites are probably unable to secure food on their own; the tibial-tarsal cleaning apparatus is underdeveloped; and, of even greater interest, the brain is reduced in size with visible degeneration in the associative centers. In the central nerve cord, ganglia 9-13 are fused into a single piece. The males are also degenerate. Their bodies, like those of the males of a few other extreme social parasites, are “pupoid,” meaning that the cuticle is thin and depigmented, actually greyish in color; the petiole and postpetiole are thick and provided with broad articulating surfaces; and the abdomen is soft and deflected downward at the tip.
In its essentials the life cycle of Teleutomyrmex schneideri resembles that of other known extreme ant parasites. Mating takes place within the host nest. The fecundated queens then either shed their wings and join the small force of egg layers within the home nest or else fly out in search of new Tetramorium nests to infest. Stumper found that the queens could be transferred readily from one Tetramorium colony to another, provided the recipient colony originated from the Saas-Fee. However, Tetramorium colonies from Luxembourg were hostile to the little parasites. Less surprisingly, ant species from the Saas-Fee other than Tetramorium caespitum always rejected the Teleutomyrmex. However, Buschinger (personal communication) has pointed out that the Saas-Fee population could be caespitum or impurum, or a mixture of both. In other words, the transfer might have been attempted across species.

From: AntWiki - Where Ant Biologists Share Their Knowledge.

Hölldobler, B. and Wilson, E. O. 1990. The Ants. Cambridge, Mass. Harvard University Press. Text used with permission of the authors.

When do we know an ant is an extreme, workerless inquiline?

From Chapter 12:

1. The worker caste is lost.
2. The queen is either replaced by an ergatogyne, or ergatogynes appear together with a continuous series of intergrades connecting them morphologically to the queens.
3. There is a tendency for multiple egg-laying queens to coexist in the same host nest.
4. The queen and male are reduced in size, often dramatically so; in some cases (for example, Teleutomyrmex schneideri, Plagiolepis ampeloni, Plagiolepis xene) the queen is actually smaller than the host worker.
5. The male becomes “pupoid”: its body is thickened, the petiole and postpetiole become much more broadly attached, the genitalia are more externally exposed when not in use, the cuticle becomes thin and depigmented, and the wings are reduced or lost. The extreme examples of this trend are displayed by Anergates atratulus, Pheidole neokohli, and Pheidole acutidens (see Figures 12-19 and 12-20).
6. There is a tendency for the nuptial flights to be curtailed, and to be replaced by mating activity among nestmates (“adelphogamy”) within or near the host nest. Dispersal of the queen afterward is very limited.
7. Probably as a consequence of the curtailment of the nuptial flight just cited, the populations of inquiline species are usually very fragmented and limited in their geographic distribution.
8. The wing venation is reduced.
9. Mouthparts are reduced, with the mandibles becoming smaller and toothless and the palps losing segments. Concomitantly, the inquilines lose the ability to feed themselves and must be sustained by liquid food regurgitated to them by the host workers.
10. Antennal segments are fused and reduced in number.
11. The occiput, or rear portion of the head, of the queen is narrowed.
12. The central nervous system is reduced in size and complexity, usually through reduction of associative centers.
13. The petiole and postpetiole are thickened, especially the latter, and the postpetiole acquires a broader attachment to the gaster.
14. A spine is formed on the lower surface of the postpetiole (the Parasitendorn of Kutter).
15. The propodeal spines (if present in the ancestral species) “melt,” that is, they thicken and often grow shorter, and their tips are blunted.
16. The cuticular sculpturing is reduced or lost altogether over most of the body; in extreme cases the body surface becomes strongly shining.
17. The exoskeleton becomes thinner and less pigmented.
18. Many of the exocrine glands are reduced or lost, a trait already described in some detail in the earlier account of Teleutomyrmex schneideri.
19. The queens become highly attractive to the host workers, which lick them frequently. This is especially true of the older, physogastric individuals, and it appears to be due to the secretion of special attractant substances which are as yet chemically unidentified.

Now, a new contribution to Myrmecological News. I'm still reading it but it looks great.

de la Mora, A., Sankovitz, M. & Purcell, J., 2020, "Ants (Hymenoptera: Formicidae) as host and intruder: recent advances and future directions in the study of exploitative strategies." Myrmecological News, Vol. 30, 53-71.

https://myrmecologicalnews.org…tegory&id=1559&Itemid=435
or
https://myrmecologicalnews.org…_printable.pdf&format=raw and https://myrmecologicalnews.org…supplement.pdf&format=raw

Teleutomyrmex.

From AmeisenWiki, on 14/03/2020, from 12:30 till 12:35:

“Teleutomyrmex.
Teleutomyrmex ist eine Gattung aus der Unterfamilie Myrmicinae. Sie umfasst nur zwei Arten, T. schneideri (Westalpen, Pyrenäen, Kasachstan) und T. kutteri (Südspanien, Sierra Nevada, bisher nur ein Volk gefunden).
Es handelt sich um arbeiterinnenlose Sozialparasiten in Nestern von Tetramorium cf. impurum. Die Tiere sind extrem selten. Auch von T. schneideri wurden seit der Entdeckung (1949 im Schweizer Wallis) bis 2006 nicht mehr als 15 Völker gefunden.
Die stark physogastrischen Königinnen hocken fast permanent auf dem Rücken der Wirtskönigin angeklammert. Ihre zahlreiche Nachkommenschaft kopuliert im Mutternest (vgl. Anergates atratulus). Wie die Ausbreitung erfolgt, ist unbekannt.
Verwandtschaftlich leitet sich Teleutomyrmex aus der Gattung Tetramorium ab.
Ergänzung 30. Aug. 2007: Die arbeiterinlose sozialparasitische Ameise Teleutomyrmex schneideri wurde nun erstmals in Spanien nachgewiesen. Zwei Gynen fanden sich in Bodenfallen im Norden der Provinz Léon auf 1.660 m Meereshöhe. Das sind ungefähr 700 km westlich des nächstgelegenen Fundortes in den französischen Pyrenäen. Der Fund unterstreicht, wie außerordentlich verstreut und isoliert die Vorkommen dieser Art sind (Espadaler & Cuesta 2006).”

From AntWeb, on 14/03/2020, from 12:35 till 12:55:

“Genus: Teleutomyrmex Kutter, 1950.
Current Valid Name:
Tetramorium.
Teleutomyrmex Kutter, 1950a: 82. Type-species: Teleutomyrmex schneideri, by original designation.
Taxonomic history:
Teleutomyrmex in Myrmicinae, Tetramoriini: Kutter, 1950a: 81 [subtribe Teleutomyrmini].
Teleutomyrmex as junior synonym of Tetramorium: Ward et al., 2015 10.1111/syen.12090: 16.
Genus Teleutomyrmex references:
Bernard, 1967a: 239 (diagnosis); Bolton, 1976: 309 (diagnosis, review of genus); Bolton, 1994: 106 (synoptic classification); Bolton, 1995a: 1053 (census); Bolton, 1995b: 403 (catalogue); Sanetra & Buschinger, 2000: 95 (phylogeny).
Distribution:
Geographic regions: Not found on any curated Geolocale/Taxon lists.”

“Species: Tetramorium inquilinum Ward et al., 2015.
Tetramorium inquilinum Ward et al., 2015 10.1111/syen.12090: 16.
Taxonomic history:
Replacement name for Tetramorium schneideri: (Kutter). [Junior secondary homonym of Tetramorium schneideri: Ward et al., 2015 10.1111/syen.12090: 16.
Distribution:
Geographic regions (According to curated Geolocale/Taxon lists):
Europe: France, Switzerland, Ukraine.
Biogeographic regions (According to curated Bioregion/Taxon lists):
Palearctic.
Specimen Habitat Summary:
Elevations: collected at 2050 m.”

“Species: Tetramorium kutteri (Tinaut, 1990).
Teleutomyrmex kutteri Tinaut, 1990b: 202, figs. 1-3 (q.m.) SPAIN. Palearctic. HOL.
Taxonomic history:
Combination in Tetramorium: Ward et al., 2015 10.1111/syen.12090: 16.
Distribution:
Geographic regions (According to curated Geolocale/Taxon lists):
Europe: Spain.
Biogeographic regions (According to curated Bioregion/Taxon lists):
Palearctic.
Specimen Habitat Summary:
Found most commonly in these microhabitats: 1 times Juniperus-Genista brushwood.
Collected most commonly using these methods: 1 times Hand.
Elevations: collected from 1660 - 2250 meters, 1955 meters average.”

“Species: Tetramorium seiferti (Kiran & Karaman, 2017).
Teleutomyrmex seiferti Kiran & Karaman, 2017: 148, figs. 3a, 4a, 5a, 6-8 (q.m.) TURKEY. Palearctic. Primary type information: Holotype (gyne) from Turkey, Artvin, Yusufeli, 3 km NW of Kınalıçam Village, N 40° 45' 36", E 41° 34' 46", 1801 m above sea level (a.s.l.), 25.VI.2013, 13 / 1592c, leg. K. Kiran, C. Karaman & V. Aksoy (in Collection of Biological Department, Trakya University, Edirne, Turkey). Paratypes: 16 gynes, 7 males from same nest as the holotype (14 gynes, 5 males in Collection of Biological Department, Trakya University, Edirne; 1 gyne, 1 male in Sofia University, Bulgaria; 1 gyne, 1 male in Senckenberg Museum of Natural History Görlitz, Germany.
Taxonomic history:
[Note: Kiran et al., 2017 PDF: 146, retain the paraphyletic genus Teleutomyrmex.].
Distribution:
Geographic regions (According to curated Geolocale/Taxon lists):
Asia: Turkey.
Biogeographic regions (According to curated Bioregion/Taxon lists):
Palearctic.
Specimen Habitat Summary:
Elevations: collected at 1801 m.”

“Species: Tetramorium buschingeri (Lapeva-Gjonova, 2017).
Teleutomyrmex buschingeri Lapeva-Gjonova, 2017: 151, figs. 3b, 4b, 5b (q.) BULGARIA. Palearctic. Primary type information: Holotype (dealated gyne) from Bulgaria, Eastern Rhodopes Mt., Chernichino Village, N 41° 35' 29.71", E 25° 50' 55.03", 640 m a.s.l., 25.IV.2012, leg. A. Lapeva-Gjonova; deposited in the museum collection of Sofia University, Bulgaria (BFUS). Paratypes (1 dealated gyne) from the same nest as the holotype, deposited in the National Museum of Natural History in Sofia (NMNHS), Bulgaria.
Taxonomic history:
[Note: Kiran et al., 2017 PDF: 146, retain the paraphyletic genus Teleutomyrmex.].
Distribution:
Geographic regions (According to curated Geolocale/Taxon lists):
Europe: Bulgaria.
Biogeographic regions (According to curated Bioregion/Taxon lists):
Palearctic.”

From AntWiki, on 14/03/2020, from 15:35 till 16:20:

“Teleutomyrmex.
This genus is not in use as it is currently considered to be a junior synonym of Tetramorium.
Nomenclature.
The following information is derived from Barry Bolton's New General Catalogue, a catalogue of the world's ants.
TELEUTOMYRMEX [junior synonym of Tetramorium].
Teleutomyrmex Kutter, 1950: 82. Type-species: Teleutomyrmex schneideri, by original designation.
Teleutomyrmex junior synonym of Tetramorium: Ward et al., 2014: 15.”

“Teleutomyrmex schneideri.
This taxon is a junior homonym and has been replaced by Tetramorium inquilinum.
Nomenclature.
The following information is derived from Barry Bolton's New General Catalogue, a catalogue of the world's ants.
schneideri. Teleutomyrmex schneideri Kutter, 1950a: 82, figs. 1-23 (q.m.) SWITZERLAND.
[Junior secondary homonym of Tetramorium schneideri Emery, 1898c: 145.]
Combination in Tetramorium: Ward, et al. 2015: 76.
Status as species: Stumper, 1951: 129; Brun, 1952: 73; Gösswald, 1953: 81; Bernard, 1967: 240 (redescription); Bolton, 1976: 309 (redescription); Kutter, 1977c: 167; Dlussky, Soyunov & Zabelin, 1990: 210; Bolton, 1995b: 403; Casevitz-Weulersse & Galkowski, 2009: 494; Borowiec, L. 2014: 170; Kiran, et al. 2017: 146.
Replacement name: Tetramorium inquilinum Ward, et al. 2015: 76.”

“Tetramorium inquilinum.
Tetramorium inquilinum is a workerless inquiline within the nests of Tetramorium caespitum and Tetramorium impurum.
Distribution.
Distribution based on Regional Taxon Lists.
Palaearctic Region: France, Iberian Peninsula, Russian Federation, Spain, Switzerland (type locality), Turkmenistan.
Biology.
As reported by Hölldobler and Wilson (1990) - This remarkable species was discovered by Heinrich Kutter (1950a) at Saas-Fee, in an isolated valley of the Swiss Alps near Zermatt. Its behavior has been studied by Stumper (1950) and Kutter (1969), its neuroanatomy by Brun (1952), and its general anatomy and histology by Gösswald (1953). A second population has been reported from near Briançon in the French Alps by Collingwood (1956) (see also Wegnez et al. 2015), a third in the French Pyrenees by Buschinger (1987c), and still others in the Spanish Sierra Nevada by Tinaut Ranera (1981). The latter was described as a separate species, Tetramorium kutteri (as Teleutomyrmex kutteri), by Tinaut Ranera (1990). Appropriately, the name Teleutomyrmex means "final ant."
The populations of T. schneideri, like those of most workerless parasitic ant species (Wilson, 1963), are small and isolated. The Swiss population appears to be limited to the eastern slope of the Saas Valley, in Juniper-Arctostaphylos woodland ranging from 1,800 to 2,300 m in elevation. The ground is covered by thick leaf litter and sprinkled with rocks of various sizes, providing, in short, an ideal environment for ants. The ant fauna is of a typically boreal European complexion, comprising the following free-living species listed in the order of their abundance (Stumper, 1950): Formica fusca, F. lugubris, Tetramorium caespitum, Leptothorax acervorum, L. tuberum, Camponotus ligniperda, Myrmica lobicornis, M. sulcinodis, Camponotus herculeanus, Formica sanguinea, F. rufibarbis, F. pressilabris, and Manica rubida. For some unexplained reason this little assemblage is extremely prone to social parasitism. Formica sanguinea is a facultative slavemaking species, preying on the other species of Formica. Doronomyrmex (= Leptothorax) pacis, a workerless parasite living with Leptothorax acervorum, was discovered by Kutter as a genus new to science in the Saas-Fee forest in 1945. In addition, Kutter and Stumper found Epimyrma (= Myrmoxenus) stumperi in nests of Leptothorax tuberum, as well as two parasitic Leptothorax, goesswaldi and kutteri, in nests of L. acervorum (Kutter, 1969).
Tetramorium inquilinum is a parasite of Tetramorium caespitum and Tetramorium impurum. Like so many other social parasites, it is phylogenetically closer to its host than to any of the other members of the ant fauna to which it belongs. In fact, it may have been derived directly from a temporarily free-living offshoot of this species, since T. caespitum and T. impurum (the host species at Briançon and in the Pyrenees) are the only nonparasitic tetramoriines known to exist at the present time through most of central Europe. It is difficult to conceive of a stage of social parasitism more advanced than that actually reached by Tetramorium inquilinum. The species occurs only in the nests of its hosts. It lacks a worker caste, and the queens contribute in no visibly productive way to the economy of the host colonies. The queens are tiny compared with most ants, especially other tetramoriines; they average only about 2.5 mm in total length. They are unique among all known social insects in being ectoparasitic. In other words, they spend much of their time riding on the backs of their hosts (Figure 12-1). The T. inquilinum queens display several striking morphological features that are correlated with this peculiar habit. The ventral surface of the gaster (the large terminal part of the body) is strongly concave, permitting the parasites to press their bodies close to those of their hosts. The tarsal claws and arolia are unusually large, permitting the parasites to secure a strong grip on the smooth chitinous body surface of the hosts. The queens have a marked tendency to grasp objects. Given a choice, they will position themselves on the top of the body of the host queen, either on the thorax or the abdomen. Deprived of the nest queen, they will then seize a virgin Tetramorium queen, or a worker, or a pupa, or even a dead queen or worker. Stumper observed a case in which six to eight T. inquilinum queens simultaneously grasped one Tetramorium queen, completely immobilizing her. The mode of feeding of T. inquilinum is not known with certainty. The adults are evidently either fed by the host workers through direct regurgitation or else share in the liquid regurgitated to the host queen. In any case, they are almost completely inactive most of the time. The T. inquilinum adults, especially the older queens, are highly attractive to the host workers, who lick them frequently. According to Gösswald, large numbers of unicellular glands are located just under the cuticle of the thorax, pedicel, and abdomen of the queens; these are associated with glandular hairs and are believed to be the source of a special attractant for the host workers. The abdomens of older T. inquilinum queens become swollen with fat body and ovarioles, as is shown in Figure 12-1. This physogastry is made possible by the fact that the intersegmental membranes are thicker and more sclerotized than is usually the case in ant queens and can therefore be stretched more. Also, the abdominal sclerites themselves are widely overlapping in the virgin queen, so that the abdomen can be distended to an unusual degree before the sclerites are pulled apart. The ovarioles increase enormously in length, discard their initial orientation, and infiltrate the entire abdomen and even the postpetiolar cavity.
From one to several physogastric queens are found in each parasitized nest, usually riding on the back of the host queen. Each lays an average of one egg every thirty seconds. The infested Tetramorium colonies are typically smaller than uninfested ones, but they still contain up to several thousand workers. The Tetramorium queens also lay eggs, and these are capable of developing into either workers or sexual forms (Buschinger, personal communication). Consequently the brood of a parasitized colony consists typically of eggs, larvae, and pupae of T. inquilinum queens and males mixed with those of Tetramorium workers.
The bodies of the T. inquilinum queens bear the mark of extensive morphological degeneration correlated with their loss of social functions. The labial and postpharyngeal glands are reduced, and the maxillary and metapleural glands are completely absent. The mandibular glands, on the other hand, are apparently normal. In addition, the queens possess a tibial gland, the function of which is unknown. The integument is thin and less pigmented and sculptured in comparison with that of Tetramorium; as a result of these reductions the queens are shining brown, an appearance that contrasts with the opaque blackish brown of their hosts. The sting and poison apparatus are reduced; the mandibles are so degenerate that the parasites are probably unable to secure food on their own; the tibial-tarsal cleaning apparatus is underdeveloped; and, of even greater interest, the brain is reduced in size with visible degeneration in the associative centers. In the central nerve cord, ganglia 9-13 are fused into a single piece. The males are also degenerate. Their bodies, like those of the males of a few other extreme social parasites, are "pupoid," meaning that the cuticle is thin and depigmented, actually greyish in color; the petiole and postpetiole are thick and provided with broad articulating surfaces; and the abdomen is soft and deflected downward at the tip.
In its essentials the life cycle of Tetramorium inquilinum resembles that of other known extreme ant parasites. Mating takes place within the host nest. The fecundated queens then either shed their wings and join the small force of egg layers within the home nest or else fly out in search of new Tetramorium nests to infest. Stumper found that the queens could be transferred readily from one Tetramorium colony to another, provided the recipient colony originated from the Saas-Fee. However, Tetramorium colonies from Luxembourg were hostile to the little parasites. Less surprisingly, ant species from the Saas-Fee other than Tetramorium caespitum always rejected the T. inquilinum. However, Buschinger (personal communication) has pointed out that the Saas-Fee population could be caespitum or impurum, or a mixture of both. In other words, the transfer might have been attempted across species.
Nomenclature.
The following information is derived from Barry Bolton's New General Catalogue, a catalogue of the world's ants.
schneideri. Teleutomyrmex schneideri Kutter, 1950a: 82, figs. 1-23 (q.m.) SWITZERLAND.
[Junior secondary homonym of Tetramorium schneideri Emery, 1898c: 145.]
Combination in Tetramorium: Ward, et al. 2015: 76.
Status as species: Stumper, 1951: 129; Brun, 1952: 73; Gösswald, 1953: 81; Bernard, 1967: 240 (redescription); Bolton, 1976: 309 (redescription); Kutter, 1977c: 167; Dlussky, Soyunov & Zabelin, 1990: 210; Bolton, 1995b: 403; Casevitz-Weulersse & Galkowski, 2009: 494; Borowiec, L. 2014: 170; Kiran, et al. 2017: 146.
Replacement name: Tetramorium inquilinum Ward, et al. 2015: 76.
inquilinum. Tetramorium inquilinum Ward, Brady, Fisher & Schultz, 2014: 16.
Replacement name for schneideri Kutter, 1950a: 82. [Junior secondary homonym of Tetramorium schneideri Emery, 1898c: 145.]”

“Tetramorium kutteri.
This species is an inquiline. Queens live in the nest of a different ant species, have no workers and are entirely dependent on their hosts for food. The queens produce eggs that are cared for and raised to maturity by the host workers.
Distribution.
Distribution based on Regional Taxon Lists.
Palaearctic Region: Iberian Peninsula, Spain (type locality).
Nomenclature.
The following information is derived from Barry Bolton's New General Catalogue, a catalogue of the world's ants.
kutteri. Teleutomyrmex kutteri Tinaut, 1990b: 202, figs. 1-3, photos. 1-2 (q.m.) SPAIN. Combination in Tetramorium: Ward et al., 2014: 16.”

“Tetramorium seiferti.
A Turkish social parasite found in the nest of a of Tetramorium species nesting under a stone in a pine forest.
Identification.
Kiran, et al. (2017) - Gynes differ from Tetramorium schneideri by larger CS, HTL / CS, PPW / CS, and DLO / CS (Tab. 1). They differ from Tetramorium kutteri by larger CS and DLO / CS, and by much longer and more erect pilosity on appendages and whole body. They differ from Tetramorium buschingeri by smaller CW / CL, larger DLO / CS, by the dorsolateral margins of the propodeum forming distinct carinae, and by much less developed microsculpture on lateral mesosomal sclerites and petiole.
Distribution.
Distribution based on Regional Taxon Lists.
Palaearctic Region: Turkey (type locality).
Biology.
Kiran, et al. (2017) - The type series was collected from a Tetramorium cf. chefketi nest located under a stone in a Pinus sylvestris L. forest older than 100 years. There are many trees about to die because of their old age, and therefore there are very large sun exposed areas on the forest ground. The ground is not inclined near the nest because the nest site is placed on the top of the hill.
The habitat is similar to that reported by Tinaut (1990) for the type locality of Teleutomyrmex kutteri.
Nomenclature.
The following information is derived from Barry Bolton's New General Catalogue, a catalogue of the world's ants.
buschingeri. Teleutomyrmex seiferti Kiran & Karaman, in Kiran, et al. 2017: 148, figs. 3a, 4a, 5a, 6-8 (q.m.) TURKEY.
[Note: Kiran, et al. 2017: 146, retain the paraphyletic genus Teleutomyrmex.]
Unless otherwise noted the text for the remainder of this section is reported from the publication that includes the original description.
Description.
Queen.
Head in full face view not much wider than long (CL / CW 0.951 - 1.000), lateral sides convex and rounding to slightly concave posterior margin, anterior part of head narrower than posterior one. Eyes protruding and small (EL / CS 0.224 - 0.250), occupying less than one quarter of lateral head side, ocelli relatively well developed, distance of posterior ocelli rather large (DLO / CS 0.244 - 0.275), anterior clypeal margin deeply concave, posterior margin broadly convex, mandibles atrophied, largely triangular, and with pointed apex. Antennal scape as long as head length (SL / CL 0.974 - 1.000), slightly surpassing posterior margin of head, funiculus 10-segmented, 3rd funicular segment slightly longer than 4th one and almost twice as long as 2nd; three apical segments forming a small club.
Pronotum narrower than head (PNW / CS 0.732 - 0.835), anterolaterally with small angles, in lateral view scutum raised over pronotum, dorsal surface of scutum and prescutum straight, scutellum higher than scutum, metanotum very small, like a tooth. Dorsolateral margins of propodeum developed as distinct carinae, the longer dorsal and the shorter declivitous profile of carinae forming an obtuse angle with a blunt tooth at the position where spines normally are based; distance of propodeal spiracle from posterior margin of propodeum twice as long as the distance from anterior margin to spiracle. Petiolar scale in profile triangular, with a nearly linear frontal and broadly convex caudal profile. Dorsal outline of postpetiole convex in lateral view. Gaster typically dorso-ventrally flattened in virgin females.
Head smooth and shiny, only genae between antennal sockets and compound eyes microreticulate. Mesosoma and petiole microalveolate. In dorsal view, postpetiole superficially microreticulate and gaster glabrous.
Head and hind tibiae with long erect hairs, scape and hind femur with dense semi-erect hairs. Dorsum of mesosoma, petiole and postpetiole with brush-like hairs, anepisternum and gaster almost bare, katepisternum with long decumbent hairs.
Clypeus, mandibles, antennae, and legs yellow. Rest of head, mesosoma, and petiole brown; postpetiole brownish yellow; gaster brownish yellow except yellow lateral sides and articulations.
Male.
In full face view head wider than long (CL / CW 0.910 - 0.935), lateral sides convex, rounding to straight posterior margin, anterior part of head slightly narrower than posterior one, eyes relatively big (EL / CS 0.233 - 0.272), located in the middle of head sides, ocelli well developed, distance of posterior ocelli rather large (DLO / CS 0.247 - 0.282), anterior margin of clypeus slightly notched medially, posterior margin broadly convex, mandibles atrophied and very small. Antennal scape short (SL / CS 0.871 - 0.907), scape hardly reaching posterior margin of head, funiculus 10-segmented, last three articles form a club.
Pronotum short and distinctly narrower than head (PNW / CS 0.762 - 0.805). With mesosoma in lateral view, mesonotum not raised as in female, its dorsum almost straight; scutellum strongly raised, metanotum lower than scutellum; dorsal surface of propodeum sloping posteriorly, slightly longer than declivitous surface and joining by an obtuse angle, and forming here a small tooth. Propodeal spiracle located close to anterior propodeal border. Dorsal part of petiole in profile almost rectangular, petiole distinctly wider than the distance between propodeal teeth, postpetiole very wide and short in dorsal view. Gaster dorsoventrally flattened.
Posterior margin of subgenital plate convex. Shape of sagitta and subgenital plate more similar to Teleutomyrmex kutteri on the contrary volsella completely different from T. kutteri and T. schneideri.
Head densely microreticulate and matt, mesosoma and petiole densely microalveolate. Postpetiole, first and second abdominal tergites with very dilute microreticulum and rest of gaster glabrous.
Long hairs absent or very rare. Short, erect and suberect hairs present on head, scape, mesosoma, petiole, and postpetiole. Gaster bare.
Lower parts of head, antennae, and legs yellow; rest of head, mesosoma, petiole, and postpetiole yellowish brown.
Gaster brownish yellow.
Type Material.
Holotype (gyne) from Turkey, Artvin, Yusufeli, 3 km NW of Kınalıçam Village, N 40° 45' 36", E 41° 34' 46", 1801 m above sea level (a.s.l.), 25.VI.2013, 13 / 1592c, leg. K. Kiran, C. Karaman & V. Aksoy (in Collection of Biological Department, Trakya University, Edirne, Turkey). Paratypes: 16 gynes, 7 males from same nest as the holotype (14 gynes, 5 males in Collection of Biological Department, Trakya University, Edirne; 1 gyne, 1 male in Sofia University, Bulgaria; 1 gyne, 1 male in Senckenberg Museum of Natural History Görlitz, Germany.
Etymology.
The species is dedicated to Dr. Bernhard Seifert for his great contributions to this study.”

“Tetramorium buschingeri.
A Bulgarian social parasite found in the nest of a of Tetramorium species nesting under a rock in a grassland.
Identification.
Kiran, et al. (2017) - Gynes differ from the other three Teleutomyrmex species by the absence of any carinae or teeth on dorsal surface of propodeum, by a very short dorsal profile of propodeum, by a much stronger developed reticulate or alveolate microsculpture covering the whole surface of lateral mesosoma and petiole and by smaller CL / CW. Furthermore, they differ from Tetramorium seiferti by a much smaller DLO / DFC (0.567 - 0.600 vs. 0.667 - 0.786). Additional differences to Tetramorium schneideri are a larger HTL / CS and a smaller EL / CS, and to Tetramorium kutteri a much smaller CL / CW, larger PW / CS and HTL / CS, and smaller ClyW / CS.
Distribution.
Distribution based on Regional Taxon Lists.
Palaearctic Region: Bulgaria (type locality).
Biology.
Kiran, et al. (2017) - Host: The two dealated queens were found on the back of a dealated queen of Tetramorium cf. chefketi together with few host workers in the nest. The host ant species belongs to the T. chefketi species complex in contrast to the hosts from the T. caespitum / impurum complex reported for Tetramorium schneideri and Tetramorium kutteri in Europe.
The nest of Tetramorium cf. chefketi with Teleutomyrmex buschingeri was found under a stone on a southern slope of a dry grassland situated in an oak forest. The habitat type is quite different from the known habitats of related species. The altitude of 640 m, where the species was sampled, is notably less than the altitudes of the localities of the other ultimate ant parasites (1600 - 2300 m) (excluding one in Turkmenistan).
Nomenclature.
The following information is derived from Barry Bolton's New General Catalogue, a catalogue of the world's ants.
buschingeri. Teleutomyrmex buschingeri Lapeva-Gjonova, in Kiran, et al. 2017: 151, figs. 3b, 4b, 5b (q.) BULGARIA.
[Note: Kiran, et al. 2017: 146, retain the paraphyletic genus Teleutomyrmex.]
Unless otherwise noted the text for the remainder of this section is reported from the publication that includes the original description.
Description.
Queen.
Head in full face view wider than long (CL / CW 0.904), its lateral sides distinctly convex, anterior part of head narrower than posterior one, eyes small (EL / CS 0.228), lateral ocelli well developed, lateral ocelli less distant from each other than in Teleutomyrmex seiferti (DLO / CS 0.219). Anterior clypeal margin deeply concave and posterior margin rather convex. Mandibles reduced and triangular, with a pointed apex. Antennal scape distinctly longer than head length, exceeding the dorsal margin of head. Funiculus 10-segmented, 2nd segment 0.043 mm, 3rd segment 0.067 mm, 4th segment 0.055 mm long, the three apical segments form a small club. Pronotum narrower than head (PNW / CS 0.747), anterolaterally with small angles. Mesonotum raised over pronotum, its dorsum straight, scutellum higher than meso notum. Dorsal surface of propodeum much shorter than declivitous surface, both surfaces form a convex outline without any traces of propodeal teeth. Propodeal spiracle located near to anterior border of propodeum. Petiolar scale in profile triangular with straight frontal and convex posterior surface. Dorsal outline of postpetiole convex in lateral view. Gaster in virgin gynes typically dorso-ventrally flattened. Hind tibiae very long (HTL / CS 0.994) distinctly longer than in Teleutomyrmex kutteri and T. schneideri.
Clypeus and frons up to the level of anterior ocellus smooth and shiny, between compound eye and antennal socket densely microreticulate. Mesosoma and petiole densely microalveolate and matt, postpetiole slightly microreticulate and more shiny. Gaster glabrous.
Head with moderately long erect hairs. Scape with semi-erect hairs. Mesosoma and petiole with brush like hairs. Postpetiole with very short brush like hairs. Gaster bare. Hind femora with long subdecumbent hairs. Tibiae with very long, erect hairs. Katepisternum and lateral portion of propodeum with very few and short decumbent hairs.
Clypeus, mandibles, antennae, and legs yellow. Head, mesosoma and petiole dark brown. Postpetiole yellowish brown, first gaster segment brown, the rest brownish yellow.
Type Material.
Holotype (dealated gyne) from Bulgaria, Eastern Rhodopes Mt., Chernichino Village, N 41° 35' 29.71", E 25° 50' 55.03", 640 m a.s.l., 25.IV.2012, leg. A. Lapeva-Gjonova; deposited in the museum collection of Sofia University, Bulgaria (BFUS). Paratypes (1 dealated gyne) from the same nest as the holotype, deposited in the National Museum of Natural History in Sofia (NMNHS), Bulgaria.
Etymology.
The species is dedicated to Prof. Alfred Buschinger who has made great contributions to the study of socially parasitic ants.”

“Key to parasitic Tetramorium species.
This queen key is based on: Kiran, K., Karaman, C., Lapeva-Gjonova, A. & Aksoy, V. 2017. Two new species of the “ultimate” parasitic ant genus Teleutomyrmex Kutter, 1950 from the Western Palaearctic. Myrmecological News 25: 145-155.
Originally assigned their own genus, Teleutomyrmex, these ants prey on other Tetramorium species. The species Tetramorium atratulum is not included in this key and bears no resemblance to the other parasites in the genus.
Males of T. buschingeri have yet to be collected.
1
Gynes . . . . . 2
Males . . . . . 5
2
Carinae or teeth on dorsal surface of propodeum absent, dorsal profile of propodeum much shorter than the declivitous one. All lateral surfaces of mesosoma and petiole covered by a well-developed reticulate or alveolate microsculpture. Head length index CL / CW < 0.945. Southern Balkans . . . . . Tetramorium buschingeri.
Carinae or teeth on dorsal surface of propodeum present, dorsal profile of propodeum not much shorter than the declivitous one. Surfaces of lateral mesosoma and petiole only in patches covered by a reticulate or alveolate microsculpture or completely smooth. Head length index CL / CW > 0.945 . . . . . 3
3
Scape long, SL / CS > 1.00. Distance of frontal carinae clearly larger than petiolar width, DFC / PW > 1.096. Size small, CW < 464 μm. Scapes and tibiae with weaker, largely decumbent pilosity. Southern Iberia . . . . . Tetramorium kutteri.
Scape shorter, SL / CS < 1.00. Distance of frontal carinae not much larger than petiolar width, DFC/PW < 1.096. Size larger, CW > 464 μm. Scapes and tibiae with profuse erect or suberect pilosity . . . . . 4
4
Ratio of distance between lateral ocelli and large diameter of complex eye larger: DLO / EL 0.93 - 1.11. Katepisternum with many long decumbent hairs, posterior corners of head posterior of the eyes smooth, absolute scape length larger: SL > 457 μm. Anatolia . . . . . Tetramorium seiferti.
Ratio of distance between lateral ocelli and large diameter of complex eye smaller: DLO / EL 0.70 - 0.80. Katepisternum without or only with a few decumbent hairs, posterior corners of head posterior of the eyes densely microreticulate, absolute scape length smaller: SL < 457 μm. Alps and Pyrenees . . . . . Tetramorium inquilinum.
5
Anterior clypeal margin straight . . . . . 6
Anterior clypeal margin concave medially . . . . . Tetramorium seiferti.
6
Subgenital plate broadly convex, sagitta with sinusoidal shape . . . . . Tetramorium kutteri.
Subgenital plate slightly concave, sagitta broadly convex . . . . . Tetramorium inquilinum.”

And the species not included in all 3 pages:

The undescribed species from Turkmenistan… See Dlussky, et al. 1990.